Austrotrema bicinctulum. 15 Austrotrema myriocarpum. Austrotrema terebrans. Nadvornikia. Key to Mazaediate Species. Key to Non-Mazaediate Species. Nadvornikia expallescens. Nadvornikia hawaiiensis. Nadvornikia peninsulae. Nadvornikia sorediata. 21 Wirthiotrema. Key to Species. Wirthiotrema desquamans. Wirthiotrema duplomarginatum. Wirthiotrema glaucopallens. Wirthiotrema santessonii. Wirthiotrema trypaneoides.
Shifts in climate along elevation gradients structure mycobiont–photobiont associations in lichens. We obtained mycobiont (lecanoroid Lecanoraceae) and photobiont (Trebouxia alga) DNA sequences from 89 lichen thalli collected in Bolivia from a ca. 4,700 m elevation gradient encompassing diverse natural communities and environmental conditions. The molecular dataset included six mycobiont loci (ITS, nrLSU, mtSSU, RPB1, RPB2, and MCM7) and two photobiont loci (ITS, rbcL); we designed new primers to amplify Lecanoraceae RPB1 and RPB2 with a nested PCR approach. Mycobionts belonged to Lecanora s.lat., Bryonora, Myriolecis, Protoparmeliopsis, the “Lecanora” polytropa group, and the “L.” saligna group. All of these clades except for Lecanora s.lat. occurred only at high elevation. No single species of Lecanoraceae was present along the entire elevation gradient, and individual clades were restricted to a subset of the gradient. Most Lecanoraceae samples represent species which have not previously been sequenced. Trebouxia clade C, which has not previously been recorded in association with species of Lecanoraceae, predominates at low- to mid-elevation sites. Photobionts from Trebouxia clade I occur at the upper extent of mid-elevation forest and at some open, high-elevation sites, while Trebouxia clades A and S dominate open habitats at high elevation. We did not find Trebouxia clade D. Several putative new species were found in Trebouxia clades A, C, and I. These included one putative species in clade A associated with Myriolecis species growing on limestone at high elevation and a novel lineage sister to the rest of clade C associated with Lecanora on bark in low-elevation grassland. Three different kinds of photobiont switching were observed, with certain mycobiont species associating with Trebouxia from different major clades, species within a major clade, or haplotypes within a species. Lecanoraceae mycobionts and Trebouxia photobionts exhibit species turnover along the elevation gradient, but with each partner having a different elevation threshold at which the community shifts completely. A phylogenetically defined sampling of a single diverse family of lichen-forming fungi may be sufficient to document regional patterns of Trebouxia diversity and distribution.
Biotic specialization holds information about the assembly, evolution, and stability of biological communities. Partner availabilities can play an important role in enabling species interactions, where uneven partner availabilities can bias estimates of biotic specialization when using phylogenetic diversity indices. It is therefore important to account for partner availability when characterizing biotic specialization using phylogenies. We developed an index, phylogenetic structure of specialization (PSS), that avoids bias from uneven partner availabilities by uncoupling the null models for interaction frequency and phylogenetic distance. We incorporate the deviation between observed and random interaction frequencies as weights into the calculation of partner phylogenetic α‐diversity. To calculate the PSS index, we then compare observed partner phylogenetic α‐diversity to a null distribution generated by randomizing phylogenetic distances among the same number of partners. PSS quantifies the phylogenetic structure (i.e., clustered, overdispersed, or random) of the partners of a focal species. We show with simulations that the PSS index is not correlated with network properties, which allows comparisons across multiple systems. We also implemented PSS on empirical networks of host–parasite, avian seed‐dispersal, lichenized fungi–cyanobacteria, and hummingbird pollination interactions. Across these systems, a large proportion of taxa interact with phylogenetically random partners according to PSS, sometimes to a larger extent than detected with an existing method that does not account for partner availability. We also found that many taxa interact with phylogenetically clustered partners, while taxa with overdispersed partners were rare. We argue that species with phylogenetically overdispersed partners have often been misinterpreted as generalists when they should be considered specialists. Our results highlight the important role of randomness in shaping interaction networks, even in highly intimate symbioses, and provide a much‐needed quantitative framework to assess the role that evolutionary history and symbiotic specialization play in shaping patterns of biodiversity. PSS is available as an R package at https://github.com/cjpardodelahoz/pss .
Geochemistry and mineralogy of rocks play important roles in the occurrence of individual lichen species and assembly of lichen communities. Whereas lichens of metal-enriched settings have been a focus of study for many decades, only a few such lichen inventories exist for North America. We reexamined the lichen biota of Pine Hill, a serpentine outcrop on Little Deer Isle, Maine and Callahan Mine, a copper-and zinc-enriched Superfund site in Brooksville, Maine by conducting additional field surveys and reexamining unidentified taxa from previous collections. To better characterize the substrates upon which the lichens were found, we conducted elemental analyses via x-ray fluorescence and inductively coupled plasma-mass spectrometry on rock samples collected at Pine Hill and recorded pH, electrical conductivity, and elemental concentrations via inductively coupled plasma mass spectrometry on soil samples from Callahan Mine. The re-investigation of lichens of the two metal-enriched sites resulted in the addition of 20 taxa to Pine Hill and 10 taxa to Callahan Mine. These include Dermatocarpon leptophyllodes, Placynthiella hyporhoda, Pyrenocarpon thelostomum, and Vezdaea acicularis, all recorded for the first time from New England. In addition, we report the first documented records since the late 19th to early 20th century for New England of Porocyphus coccodes, Sarcosagium campestre, and Steinia geophana, and the first such record for Maine for Coccocarpia palmicola. Stereocaulon condensatum and S. subcoralloides, both considered as rare in New England, were also collected from Callahan Mine.
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